Specialist or refugee: Microhabitat use and competition between two sympatric woodland salamanders

Co-occurrence between species may be mediated by ecological differences or competitive interactions, but the nature of these interactions can vary across spatial scales. At coarse scales, species may appear to broadly co-occur, but at fine scales, particularly for species with small home ranges, species may utilize different aspects of the microhabitat such that co-occurrence is limited and competitive interactions are reduced or eliminated. We investigated the microhabitat use of two morphologically and ecologically similar woodland salamanders (the Red-backed Salamander, Plethodon cinereus, and the Northern Ravine Salamander, P. electromorphus) that are distributed throughout the Alleghany Plateau in eastern North America to determine whether they partition habitat at finer scales. We also tested if P. electromorphus is a specialist on habitat types that allow it to exclude P. cinereus, or if P. electromorphus uses lower quality habitat as a refuge to avoid P. cinereus. At a site of microsympatry, we repeatedly sampled plots during two seasons, each within a different area of the landscape, and used occupancy and abundance modeling to assess the extent of co-occurrence and test whether the species use different microhabitat conditions. Plethodon cinereus greatly outnumbered P. electromorphus and occupied a greater proportion of plots, most plots where we detected P. electromorphus we also detected P. cinereus. In contrast to patterns documented in some montane species pairs, in which a high-elevation habitat specialist excludes a low-elevation generalist, we found that P. electromorphus primarily used microhabitat conditions favorable for both species, such as north-facing slopes, yet did not have a negative effect on the abundances of P. cinereus. In the presence of an abundant competitor, microhabitat use of the less abundant species may narrow toward optimal conditions with sufficient resources to facilitate co-occurrence.